"So, naturalists observe, a flea has smaller fleas that on him prey; and these have smaller still to bite ’em; and so proceed ad infinitum."
- Jonathan Swift

July 11, 2014

Special Report: #ASP2014 (Australia) Part I: The Wild World of Parasites

Photo by Lisa Jone
Recently I attended the annual meeting of the Australian Society for Parasitology (ASP) - it also happened to be the 50th anniversary of the Society, so it was kind of a big deal for the ASP. The first day featured an opening speech by Australian Chief Scientist Ian Chubb. In it, he discussed the many people of the world of dying and suffering from preventable infectious diseases which is the price of poverty, poor sanitation and ignorance. He also talked about how the political priorities of Australia's current government does a great disservice to science, and the lack of long range strategies regarding science, technology, and engineering is holding back Australia as a nation.

He likened it to scattering pieces of a jigsaw puzzle with no means of connecting them, and it is detrimental to Australia's future. Chubb also emphasised that science is vital to the future of Australia and the importance of engaging the public and the next generation with the importance and awe of science (which I hope that I am playing at least a tiny part in by writing this blog!). Speaking of science, as that is what you came to this blog for after all, what kind of parasitology research caught my attention at the conference? For this post I will mostly discuss the presentation on wildlife parasites I saw at the conference.

There was a very interesting plenary talk by Vanessa Ezenwa about how multiple parasites infecting the same animal can influence the resulting pathology inflicted by those parasites upon the host. She presented a case in African buffaloes whereby the removal of parasitic worms affected the disease severity of bovine tuberculosis (bovine TB). There appears to be a trade-off between being resistance to macroparasites (worms) and microparasites (TB bacteria), with buffaloes that are more resistant intestinal worms being less able to mount a response to invasion by the tuberculosis bacteria. It seems as if the worms are pre-occupying the host immune budget, thus allowing the TB bacteria to slip by. However, if the buffaloes are treated with anti-parasite drugs that rid them of their worms, they were able to stop the TB bacteria dead in their tracks. Who would have thought treating buffaloes for their worm infections would also rid them of TB? Ezenwa's study shows the importance of considering the entire parasite community of a host animal and taking an ecological approach to considering host-parasite interactions.

On the subject of ecology, Haylee Weaver presented a talk based on a project that we have been collaborating on regarding parasites that infects animals with semelparous life-cycle - like the Sockeye Salmon, or the Antechinus - better known as the the little Australian marsupial that "has so much sex it disintegrates" followed by a talk I gave on a comparative analysis study I conducted on with Janet Koprivnikar which compared the nematodes fauna of migratory and non-migratory birds.
Photo of sea lion family by DaveDiver from Wikipedia

Jan Šlapeta presented research into a species of hookworm in Australian Sea Lions (Neophoca cinerea). This parasite - Uncinaria sanguinis - exploits the dependency between mother and offspring. The hookworm lives in female sea lions but unlike other hookworms, it does not lay eggs which are passed out in the host's fece, instead it is transferred to the pup via the mother's milk - only then does the worm mature into an egg-laying adult stage like other hookworms. Because of this transmammary transmission, male sea lions are considered to be a dead-end host for U. sangunis.

Because female sea lions do not tend to dispersed, it would be expected that the population of the parasite would be highly structured, but Šlapeta and colleagues found that was not the case, and that the population genetics of U. sanguinis is not as well segregated as expected. This raise many questions about the ecology of this parasites, such as whether other species of sea lions and seals serve as alternative hosts? Or perhaps the males are not dead-end hosts after all? Or can crustaceans like shrimps act as paratenic (transport) hosts for the parasite?

Elsewhere at the conference, there were many posters and talks on Cryptosporidium which seems to be a popular topic of research among Australian parasitologists. There are many different species of Cryptosporidium and not all of them infect humans - though some have potential to jump from their usual hosts into humans. For example, Australian marsupials and multitude of other wildlife are host to various species of Cryptosporidium and Michelle Powers presented a talk on the current state of knowledge about this genus of parasite and concluded there are still many different host species that can be harbouring undescribed species of Cryptosporidium.

Mammal ectoparasites were also also featured at the conference, with a poster presentation by Clare Anstead on the specificity of ticks that infects small mammals as well as their bacterial communities - just as there are generalists ticks that feed from a variety of host species and more picky specialists that stick to just one or two, it seems that the same goes for their bacterial occupants in regards to the species of ticks they inhabit. Speaking of ticks, Stephen Barker announced the launch of a 140 page monograph he and Alan Walker wrote on the ticks of Australia which are found on domesticated animals and humans. And it is available for free for all to download here, which I am sure will tickle the fancies of all tick fans.
Photo of crocodile farm by Cecil Lee
Moving on from parasites of furry hosts to more scaly ones, Simon Reid presented an unusual case of parasitism on a crocodile farm. We have featured various crocodilian parasites on this blog before, in this case these crocodiles on the farm end up being infected with a muscle-burrowing worm due to human action.

The practice of raising crocodilians in a farm setting has come about due to the demand for crocodilian skin product, but another product of such farms is crocodile meat. Since the meat is meant for human consumption, this has led to them being tested for parasites and pathogens, which in turn led to the discovery of an unexpected species parasitic worm in the muscles - Trichinella papuae. Trichinella is also known as "the worm that would be a virus" and normally, crocodiles are known to be infected by their own species of Trichinella - Trichinella zimbawensis. But T. papuae is normally a pig parasites - so how did they end up in a crocodile? Well the obvious answer is that those crocodiles were being fed with pigs - but it also provides an interesting insight into the biology of the parasite itself because their presence in crocodile muscles means that even though  T. papuae normally dwell in an mammal, it is also adaptable enough that it can also survive in a host with a rather different physiology to its usual host.

Speaking of scaly hosts, fish are the most diverse vertebrate animal on the planet and talks about their parasites had a considerable presence at this conference. In Part Two of my special report on ASP 2014, I will be covering fish parasites - including how to make invisible parasites visible, what is the relationship between tongue-biters and face-huggers, and what parasite you might find in the fins of an epaulette shark. All that and more will be revealed in my next post on ASP 2014.

June 25, 2014

Ismaila belciki

Photo of infected Janolus fuscus
used with permission from Jeff Goddard
If you ever find yourself down by the sea, you may come across some very flamboyant sea slugs call nudibranchs. But beneath their colourful exterior, some of them are harbouring a dark secret in the form of a very strange looking parasite. These parasites live hidden inside the main body cavity of their molluscan host, so if you are unfamiliar with this particular critter, you might not even notice it. The main thing that gives away their presence are a pair of egg sacs poking out of the sea slug (see photo on the right). Those egg sacs belong to a parasite call Ismaila belciki - it is a crustacean, though it looks more like one of Cthulhu's lovechild or something out of Men In Black.

Ismaila and other copepods of the Splanchnotrophidae family are specialist parasites of sea slugs and they can get pretty big in comparison with their host, taking up substantial room and resources. Ismaila belciki infects Janolus fuscus, a nudibranch found along the west coast of North America from Alaska to California, as well as the shores of northern Japan. In some areas, such as Coos Bay, Oregon where the study we are featuring today took place, up to 80% of the slugs are infected with this odd creature. Having such a big parasite sitting in the middle of slug's body soaking up nutrient obviously carries some kind of cost - but just how much?

Photo of a female Ismaila belciki with an
embraced dwarf male front and centre.
Photo by and used with permission
from Maya Wolf
A pair of researchers from University of Oregon decided to find out just how costly this parasite is to its host. They compared the growth, survival, and reproductive capacity of infected and uninfected J. fuscus, and measured how much resources the parasite takes up.

While I. belciki did not seem to interfere with sea slug's growth, infected slugs do have a lower survival rate. Additionally, they have shrunken gonads that are only capable of producing about half as many eggs as healthy slugs. But the reproductive capacity of those afflicted sea slug suffers not just in terms of quantity, but in quality as well. In addition to producing fewer eggs, infected slugs also produced eggs that were smaller, and the baby slugs that hatch out of them also have lower survival rates.

So it seems I. belciki can be very harmful indeed, but it cause even greater harm if the parasite itself is breeding. The researchers noted that I. belciki bearing developing egg sacs exert a greater toll on the host than egg-free parasites. A female I. belciki is an egg-laying machine that can churn out over 88000 embryos per month and all the expenses for that are paid for by the host. To fuel the development of its eggs, I. belciki draws from the same pool of resources that the host normally use for its own egg production. Slugs with brooding I. belciki produce even fewer eggs than those that are "just" stuck with an egg-free parasite.

It is as if the sea slug is a factory that has been retooled from solely making slug babies into one which now has to divert some of its attention and raw material to making parasite babies too, via a proxy in the form of a female I. belciki. Given that Janolus fuscus usually only live for five months, by shorten their lives and severely reducing their reproductive capacity, I. belciki might actually be putting a natural check on the population growth of these flamboyant nudibranchs.

Wolf, M., & Young, C. M. (2014). Impacts of an endoparasitic copepod, Ismaila belciki, on the reproduction, growth and survivorship of its nudibranch host, Janolus fuscus. International Journal for Parasitology 44: 391-401.

P.S. I will be attending the annual Australian Society for Parasitology annual conference in Canberra, Australia between 30th June to 3rd July. So watch for tweets about highlights from conference at my Twitter @The_Episiarch! Meanwhile, I have written a article for The Conversation about the crab-castrating barnacle Sacculina carcini - you can read it here.

June 10, 2014

Anilocra nemipteri

Photo from Figure 1 of the paper
The parasite in the study being featured today makes a living riding around on the top of a fish's head and occasionally gnawing on its face. It is in the same family as the infamous tongue-biter, the Cymothoidae, though technically this one is more of a face-hugger.

Anilocra nemipteri is found on the Great Barrier Reef of Australia and it makes a living by hitching a ride (and feeds) from the bridled monocle bream, Scolopsis bilineata. It is a pretty common parasite - in some areas, up to 30 percent of monocle bream carry one of these crustaceans on their head like a nasty blood-sucking beret that stay attached for years.

As you can see from the photo, A. nemipteri is a fairly big parasites comparing with the size of the fish (in some case they can reach as almost one-third the length of the host fish!), and having a parasite of that size hanging off your face is going to be quite a drag - literally. That is bad news for a little fish like the monocle bream that needs to make a quick getaway from any hungry predators on the reef. So just how much of a drag is A. nemipteri? A related species - Anilocra apogonae - which clings to the cardinal fish (Cheilodipterus quinquelineatus) is known to cause their host to swim slower and have lower endurance. Does the same apply for A. nemipteri and the monocle bream?

To find out, scientists compared how quickly the fish can respond to an attack and their Flight Initiation Distance (FID) in both a laboratory setting and in the field. The FID is the distance from a predator at which an animal decides to flee - risk-takers have a shorter FID. They divided the monocle bream into three different groups: parasite-free fish, fish carrying an A. nemipteri, infected fish which just had their parasite removed.

Photo from Figure 1 of the paper
The research team simulated an attack by a bird (with a weighted PVC pipe) on fish in specially-designed experimental tanks and filmed the response to measure the fish's reaction time to the attack. Even though one would think all that face-gnawing from A. nemipteri would have weakened their host, and not to mention the body of the parasite itself causing significant drag, the escape performance of parasitised fish was not all that different from unparasitised - they reacted and got away from the attack just as quickly as their unburdened buddies. In the field experiment, the scientists donned snorkelling gear and tried to approach any monocle breams they spotted and measured how close they could get to the fish before they fled. There, they found parasitised fish have a slightly shorter FID than parasite-free fish, but not significantly so.

Fish that are infected by A. nemipteri are smaller than uninfected ones, and it just so happen that smaller fish tend to allow predators to get closer to them before fleeing. But whether this is due to the parasite is another matter. Are parasitised fish smaller because their growth have been stunt by A. nemipteri? Or does this face-hugger simply prefer smaller fish because larger and older fish might have built up an immunity to it?

Though it may seem less exciting when we find a parasite doesn't cause much behavioural changes in its host, it is vital to our understanding of host-parasite relationships. Perhaps it means the host is able to compensate for the presence of the parasite. Also it is not clear what the long term cost of having A. nemipteri might be over the life time of the fish. It is also important to treat such a case in its context. Unlike other parasite which have a life-cycle and depend upon its host getting eaten by a predator to reach maturity, A. nemipteri is an external parasite that simply sticks to a host and stay for life - if the parasitised fish is eaten by a predator, it'll go down with the host like a bit of garnish and be digested too.

So it is probably just as well that A. nemipteri is not too much of a drag to have around.

Binning, S. A., Barnes, J. I., Davies, J. N., Backwell, P. R., Keogh, J. S., & Roche, D. G. (2014). Ectoparasites modify escape behaviour, but not performance, in a coral reef fish. Animal Behaviour 93: 1-7.

May 25, 2014

Loxothylacus panopei

Photo by Inken Kruse via the Hare Lab
Some parasites can manipulate their host's behaviour in very spectacular ways, but there are also other parasites that change their host's habits in more subtle manners. While such alteration to the host can seem fairly minor, they can still result in some very profound impact on the rest of the ecosystem.

There is a group of parasitic barnacles call Rhizocephala (the most well-known species is Sacculina carcini) that are capable of castrating their host, turning them into unwitting babysitters that nurture the parasites' brood. The infected crab display some very obvious changes to their behaviour, and in some cases, their appearance. But the study we are featuring today shows that apart from turning them into doting mothers for the parasite's babies, these barnacles can also alter the crab's behaviour in less obvious ways that have ramifications for other marine inhabitants.

The flatback mud crab (Eurypanopeus depressus) lives in estuaries on the coast of South Carolina and it is infected by a species of rhizocephalan call Loxothylacus panopei. In addition to doing the usual host castrating and commandeering trick, L. panopei also changes how this crab responds to potential prey. Usually, the mud crab has an omnivorous diet, dining on algae as well as worms, smaller crustaceans, and sponges. Sometimes they may also have a crack at more armoured prey like mussels. But crabs that are infected with L. panopei lose their appetite for such shell-covered fares.

When researchers offered uninfected crabs with piles of mussels, the crabs acted like they were at an all-you-can-eat seafood buffet and ate as much as they can - the more mussels the researchers presented them with, the more they ate. But no matter how many mussels they offered to crabs that were infected with L. panopei, they simply eat one and call it a day. The parasitised crabs also took longer to get their act together and this seems to be related to the size of the crab's parasite - the larger the parasite has grown, the longer the crab takes to start digging into a mussel.

Based on a field survey of the estuary where the study took place, the researcher concluded that about a fifth of the crab at that location were infected with L. panopei. Given the effects that L. panopei has on their crab's appetite for shellfish, it seems that the mussels might have an unlikely ally in the form a parasitic barnacle. The finding of this study share some parallel to another paper that we featured on this blog earlier this year, on the muscle-wasting parasite that infects a predatory shrimp and curb its otherwise ravenous appetite.

Ecosystems are made up of complicated networks of biological interactions and parasites can mediate predator-prey interactions in different, and sometimes conflicting ways. While some parasites can make prey animals more vulnerable or accessible to predators, there are other like L. panopei that may be reducing the appetite of the said predators. The subtle interplay of such parasite-mediated interactions are often overlooked or ignored, but their effects on the ecosystem are certainly there if you know what to look for.

Toscano, B. J., Newsome, B., & Griffen, B. D. (2014). Parasite modification of predator functional response. Oecologia 175: 345-352.

May 8, 2014

Nematocenator marisprofundi

Parasitism is the most common mode of life on Earth and it can found everywhere, in all kinds of environments. Even in extreme places such deep sea hydrothermal vents, amidst hellish geysers pouring out hot sulfide or seeping methane, parasitism carries on as usual - the players may change, but the game stays the same. While on this blog most of the nematodes we have featured are the parasites, in this particular case, they play the role of the host.

SEM photo of D. marci from the paper
Laying about 85 kilometres off the coast of Oregon, under about half a mile (800 metres) of water is the Hydrate Ridge methane seeps. These vents are covered in mats of sulfide oxidizing bacteria which are crawling with worms - mostly nematodes from the genus Desmodora. One of these species happens to be a host to the parasite we are featuring today - Nematocenator marsiprofundi - which translates into "nematode eater of the deep sea". It is a microsporidian - a group of single-cell parasite somewhat related to fungi, and taxonomically speaking, N. marsiprofundi lies right near the base of the split between these two groups.

Microsporidians are found in a wide range of animals including vertebrates such as fish and reptiles, as well as invertebrates such as insect, crustaceans, and nematode worms. The host of N. marsiprofundi, a nematode named Desmodora marci (see above), is one of the more abundant animal at methane seeps. There can be as many as twenty worms for each millilitre of carbonate rocks from such locales, and over half of those worms would be infected with N. marsiprofundi. This parasite seems to be common at such vents and were found at sites which are 15 kilometres, so N. marsiprofundi is not localised to just a particular location and/or worm population.

Spores of N. marisprofundi
(image from the paper)
The spores of this parasite (see left) are mostly found in the worm's reproductive tract; in female worms, the spores sit in the uterus next to the eggs, and in the male, the spores lined the worm's sperm duct and cloaca. This led the researchers who found this parasite to suggest that N. marisprofundi is sexually transmitted between its host. However, they also noticed some stages of the parasite were situated in the body wall, where they seem to degrade and digest the worm's muscle tissue, not unlike the microsporidian we featured a two months ago which infects an amphipod that has become invasive in Central Europe.

While their presence in the body wall and the effects they have on their host's muscle indicates they can be quite harmful and may transmit through means other than the worms' sexual activities, that is not to say that this parasite might not exploit multiple mode of transmission. Some parasite change their shape and infect different host tissue at different stages of their lives, and it is possible that N. marisprofundi can both be sexually transmitted and also eventually kill their host to allow their spores to disperse from a rotting cadaver

Studies like this shows parasites might be more common in the deep sea that we might have previously suspected, and that even in seemingly extreme environments like hydrothermal vents, there is good living to had as a parasite. Parasitism is everywhere on this planet, and while many people may think parasites are odd freaks of nature, in reality they are just a normal part of life on Earth.

Sapir, A., Dillman, A. R., Connon, S. A., Grupe, B. M., Ingels, J., Mundo-Ocampo, M., Levin, L. A., Baldwin, J. G., Orphan, V. J. & Sternberg, P. W. (2014). Microsporidia-nematode associations in methane seeps reveal basal fungal parasitism in the deep sea. Frontiers in Microbiology 5: 43.

April 20, 2014

Controrchis sp.

Extreme weather events can cause significant changes to ecosystems and their inhabitants. When Hurricane Iris made landfall at Belize, it caused widespread devastation in its wake. The study we are featuring today was a part of a larger project to look at how Hurricane Iris affected a population of black howler monkeys (Alouatta pigra) which has been monitored since 1998. In the aftermath of the hurricane, the number of monkeys in the forest decreased by 78 percent until their population began to stablise and increase three and a half years later. But aside from such outwardly visible impacts, there were also other changes afoot within the monkeys themselves.
Photo of black howler monkey by Ian Morton

A team of scientists interested in monitoring the recovery carried out a study to see how this has affected the monkeys' parasites. It is possible that these primates are harbouring higher parasite loads than they did before the hurricane due to the stress of living in a disturbed habitat. The scientists collected samples of monkey fece over the course of 3 years and look for parasite eggs. They also measured the level of cortisol, a hormone associated with stress, present in the fecal sample, and collected data on other aspects of the monkey's behaviour to see if they were associated with their parasite burden.

Photo of Controrchis eggs
from here
The black howler monkeys were found to have five species of roundworms and a species of digenean fluke (based on the presence of their eggs in the monkey poop), but the prevalence and abundance of those parasites were not associated with the level of stress hormones. Instead, nematode (roundworms) prevalence was heavily dependent on population density and the size of the groups in which the monkeys gathered. This is to be expected as these worms are transmitted via accidental ingestion of eggs or larvae from the host's feces. The more monkeys there are around in a given area, the more opportunities for these particular parasites to be passed on. This is similar to what has previously found in other studies on primate parasites. But the only factor that successfully predicted the occurrence of the digenean trematode fluke Controrchis was the amount of leaves the monkeys consumed.

While black howler monkeys usually prefer a diet filled with fruit, in the aftermath of Hurricane Iris there were no fruit-bearing plants in the forest for 18 months. So the monkeys were forced to go on a leaf-based diet instead of the fruit-based one they enjoyed before the hurricane, and the plant most readily available and palatable to the monkeys was Cecropia. These fast-growing leafy plants usually happens to be the first on the scene in the wake of such habitat disturbances. They do not contain as much fibre as other plants and have little in the way of noxious defensive chemicals - which makes Cecropia excellent fodder for the black howler monkeys. Cecropia also contains a lot of what these monkeys need in a balanced diet, so in the absence of fruits, the howler monkeys munched readily on these nutritious greens

But why is the consumption of Cecropia associated with the prevalence of Controrchis? The fluke does not use leaves and vegetation as a mean of transmission (unlike Fasciola the liver fluke), instead, Controrchis uses ants as a go-between to get in their vertebrate host. But these monkeys don't really have a taste for ants, so why is Controrchis prevalence linked to the amount of leaves they have consumed? That is because Cecropia also happens to be myrmecophtyes, or ant-plants. Monkeys that are chowing down those leafy greens are also inadvertently swallowing a lot of ants, which means taking onboard a lot of Controrchis waiting to make a connection with a suitable monkey host.

For another more detailed take on this paper, from the lead author herself, see this post here

Behie, A. M., Kutz, S., & Pavelka, M. S. (2014). Cascading Effects of Climate Change: Do Hurricane‐damaged Forests Increase Risk of Exposure to Parasites?. Biotropica 46: 25-31.

April 9, 2014

Bivitellobilharzia nairi

A little over a year ago, I wrote a post about Bivitellobilharzia loxodontae - a species of blood fluke that lives in the African forest elephant. Today I am writing about a study on another species from that genus - Bivitellobilharzia nairi - which parasitises the Indian elephant. However in a newly published study, it turns out the Indian elephant is not the only thick-skinned mammal that harbours this fluke.

Photo of Indian rhino by Krish Dulal
The study we are featuring today took place in southern Nepal at the Chitwan National Park (CNP). Researchers collected fecal samples from both domesticated and wild Indian elephants for examination and as expected, they found B. nairi eggs amongst the samples. But it was when they started looking in the poop of Indian rhinoceros that they found the unexpected. These rhinoceros do not take a dump just anywhere; they are creatures of habit and defecate at specific spots call faecal middens - which is how they mark their territory. When the researchers dug through the contents of those middens, they found blood fluke eggs amidst the rhino dung in half of the fourteen middens they sampled from.

The eggs had the characteristic look of schistosome eggs - an ovoid with a hook at one end (see below). But they were not just any blood fluke eggs, they looked very similar to the eggs of B. nairi - the elephant blood fluke. When the researchers sequenced specific marker section of the fluke eggs' DNA, they found that it matched the known sequences for B. nairi, showing that what is usually thought of as just an elephant parasite can also find a home in the Indian rhinoceros. Furthermore, the B. nairi eggs they recovered from the rhino dung were completely viable, showing that the rhino is a natural and commonly used host for this parasite and that they did not end up there by accident

Image of Bivitellobilharzia nairi egg from here
Evolutionarily speaking, elephants and rhinoceros are fairly far apart on the mammalian tree - the last common ancestor they shared lived about 100 million years ago in the era of non-avian dinosaurs. So what is an elephant schistosome doing in a rhino? Despite their specialised adaptations for living in the circulatory system and evading the immune reactions of their particular hosts, throughout their evolutionary history, schistosome have made a number of leaps across divergent animal taxa. One such jumps had allowed the ancestors of schistosomes to evolve from a sea turtle-infecting parasite into one which live in the blood of warm-blooded animals like birds and mammals. While elephants and rhinoceros have had disparate evolutionary paths for at least a hundred million years, clearly their physiology are similar enough for B. nairi to successfully survive in both. In addition, their shared habitat provided the fluke with plenty of opportunity to encounter and adapt to the rhinoceros.

So there is more than one way for two (or more) different species to end up with the same parasite. You can either share a recent common ancestry, or you can share the same habitat which gives the parasite ample opportunities to cross the evolutionary gulf between different hosts.

Devkota, R., Brant, S.V., Thapa, A. & Loker, E.S. (2014) Sharing schistosomes: the elephant schistosome Bivitellobilharzia nairi also infects the greater one-horned rhinoceros (Rhinoceros unicornis) in Chitwan National Park, Nepal. Journal of Helminthology 88: 32–40

March 26, 2014

Octopicola superba

When it comes to reproduction, most living things can be classified along a scale. At one end, you have the r-strategists (many insects and molluscs) that produce a prodigious number of offspring but few survive to adulthood. And on the other end are the K-strategists that produces only a few progeny, but to invest a lot of resources into each to ensure they are more likely to reach maturity (for example, elephants, humans, etc).

SEM photo of female
Octopicola superba from here
There is a cost/benefit trade-off inherent with being on either side of the scale because as a r-strategist, you might be producing a lot of progeny, but most of them will probably die before they get to reproduce themselves. While on the K-strategist end, by investing so much resources into each individual young, you can only afford to produce a few of them. The reproductive strategy of different organisms all fall somewhere along that continuum between low quality mass production or high quality but infrequent output, and different circumstances call for different strategies.

Textbook often use parasites as key examples of r-strategists, as a model of organisms that producing prodigious number of offspring. Indeed some internal parasites are well-known for their reproductive capacity - for example, the female blood fluke Schistosoma mansoni produces 300 to 3000 eggs per day, while tapeworms like Diphyllobothrium dendriticum can produce tens of millions of eggs per day. But not all parasites opt for quantity over quality.

The study we are featuring today examined the reproductive capacity of the parasitic copepod Octopicola superba, which, as its name indicates, lives in the common octopus. As far as a parasite goes, this crustacean seems rather innocuous and does not really cause much harm to its host. Octopicola superba can be found all over the body of the octopus but most of them are located on the skin and gills. Even though it is a parasite, it has a reproductive strategy which brings it closer to being a K-strategist.

Each female O. superba produces a clutch of only a few dozen eggs per season; if a female was to produce more than about forty eggs in a clutch, she starts reaching the upper limit of her reproductive capacity and the size of each egg (which reflects how much resources is invested into it) begins to shrunk as the brood imposes too much of a drain. This reproductive capacity varies considerably between individual; the most productive copepods are able to produce over twice as many eggs as the least productive ones, while some produced eggs that were almost twice as big as those produced by others.

Octopicola superba's reproductive strategy also shifts during different seasons; in winter, they produced a larger clutch of smaller eggs, whereas in summer they produce a smaller clutch of bigger eggs. Such season shifts has been observed in other parasitic copepods, though for O. superba, the reason for them doing so remains unknown. Despite these seasonal and individual differences, overall O. superba is certainly low-key when it comes to reproduction - even the most fecund female had just above sixty eggs in a clutch and the rest mostly produced between thirty to forty eggs.

So why has this parasitic copepod evolved to produce so few eggs compared with parasites like tapeworms and blood flukes that pump out thousands or even millions of eggs on a daily basis? It might have something to do with the habits of its host.

Octopus tend to be territorial homebodies that likes to stay in their little corner of the sea. Previous analyses indicate that hosts with such sedentary habits tend to select for parasitic copepods that produce larger eggs. Unlike one infecting more mobile animal (like a fish), parasites of sedentary animals cannot rely upon their host's routine daily movement to bring them into contact with new hosts. Therefore, they must do so under their own steam. By investing more into each egg, the female O.superba ensures each of her babies are better equipped for the long journey to find a new home, even if it means she can only produce just a few dozen of them at a time.

With offspring, you can only invest so much into them - at some point, they are on their own

Cavaleiro, F. I., & Santos, M. J. (2014). Egg number-egg size: an important trade-off in parasite life history strategies. International Journal for Parasitology 44:173-182

March 9, 2014

Cucumispora dikerogammari

Invasive species can be very disruptive - cane toads, rabbits, water hyacinth, and zebra mussels are just a few well-known examples of species that have been introduced to areas outside of their original geographic range and have caused extensive ecological disruption in their new home. One of the hypotheses for why some introduced species become so successful when they arrive at a new region is called the "enemy release hypothesis". In their new home, introduced species run amok as they are no longer hounded by their usual foes that would otherwise keep their population in check.
Top: A heavily infected amphipod
Bottom: Spores of C. dikerogammari
Photo from here

Dikerogammarus villosus is an amphipod (a little, shrimp-like crustacean) from the Ponto-Caspian that has invaded western and central Europe, and is now also found in the United Kingdom. They might only grow up to a little over an inch long, but they are voracious little predators that eat everything smaller than themselves, including each other. Released from their usual predators and parasites, D. villosus rips through the freshwater life of its new neighbourhood. But they have not completely escaped from their past foes; one parasite has managed to come along for the ride, and it is a microsporidian called Cucumispora dikerogammari.

As far as the parasite goes, Cucumispora dikerogammari is a pretty nasty one. It invades the host's muscles, reproduces prolifically and eventually kills the host by overwhelming it with sheer numbers. There is some concern that this parasite can spill over into the native invertebrates and add insult to injury to the local stream life. But on another hand, a new study shows that this parasite might be one of the few things holding back this voracious invasive amphipod from causing even more destruction.

A group of scientists from France conducted a study to looked at how C. dikerogammari affects the activity levels and appetite of D. villosus. They observed the behaviour of both infected and uninfected amhipods in a water-filled glass tube and noticed that amphipods at a late stage of infection that are visibly "filled to the brim" with parasite spores are actually more active than healthy amphipods or those that are not visibly parasitised because they are at a much earlier stage of the infection.

Close-up of a C. dikerogammari spore from here
Furthermore, they also presented amphipods with midge larvae (also known to some as "bloodworms") to see how many they ate. Both infected and uninfected D. villosus pounced on those insect larvae, but the heavily infected amphipods ate far less than the healthy ones. For whatever reason, this parasite seems to cause D. villosus to lose its appetite, and given this crustacean's reputation of eating everything that it can get its claws around, this may have significant ecological ramifications. It could mean that C. dikerogammari may be subtly reducing the impact these amphipods have on the areas where they have been introduced.

But why would heavily-infected D. villosus, which would have much of their muscle mass already converted to parasite spores by C. dikerogammari, be more active? Well, it could just be an odd manifestation of the disease, but if it is, it is certainly a useful one for this parasite - as it depends upon cannibalism for transmission to new hosts. Dikerogammarus villosus are rather homely creatures and usually prefer to stay under a shelter and wait for potential prey to wander by. By getting their host out and about, C. dikerogammari might increase the chances that its host will either run into one of its cannibalistic buddies, or die out in the open where it can be scavenged by other D. villosus.

It seems that for this little invasive amphipod, no matter how far you go, you can never really run away from your past (foes).

Bacela-Spychalska, K., Rigaud, T., & Wattier, R. A. (2013). A co-invasive microsporidian parasite that reduces the predatory behaviour of its host Dikerogammarus villosus (Crustacea, Amphipoda). Parasitology 141: 254-258.

February 14, 2014

Gordionus chinensis

Hairworms are known for their ability to make their host go for an impromptu (and terminal) swim in a stream or a pond, but by doing that they are not just sending ripples through the water, but also into the surrounding ecosystem. The paper we are looking at today features a species of hairworm from Japan call Gordionus chinensis - this parasite infects three different species of forest-dwelling camel crickets from the genus Diestrammena.

Photo by Danue Sachiko from here
The scientists who conducted the study that this paper is based on wanted to find out what happens to the the cricket population and their hairworm parasites after their home forest has been cut down. They conducted an observational field study at an experimental forest in the upper parts of the Totsu River at Nara Prefecture, Japan. The forest was originally clear-cut in 1912 and 1916 and since then, parts of it have been replanted and cut down at different point in time over the last century. Each study site corresponds with a different replanted forests of Japanese cypress ranging from 3 to 48 years old.

These scientists found that the camel crickets began returning a few years after a forest has been replanted, their abundance steadily increasing and eventually reaching a peak after the forest has been standing for at least 30 years. But their hairworm parasites did not return with similar gusto. In fact, they estimated that only second-growth forests that are more than 50 years old have hairworm populations that are as abundance as those found at undisturbed sites.

One possible reason for the hairworms' slow recovery is their complex life cycle which requires infection of more than one host. The replanted forest might be lacking some of the other host G. chinensis needs to complete its life cycle. Because parasites has such a negative public image, a forest which is free of parasites (or at least a specific parasite) might sound good to most people. But these hairworms actually play a very vital role in the ecosystem.

By causing their cricket host to jump into a stream, they actually serve as a kind of fast food delivery service for the fish living in those streams. A cricket infected with a hair worm is 20 times more likely to stumble into a stream and become fish food than an uninfected cricket - so fish which would not usually get to feed on such large land-loving insects on a regular basis can now do so thanks to the hairworm, and it has calculated that this straight-to-your-stream food delivery service accounts for 60% of the trout population's energy intake in some watersheds.

For hairworms, new forests just do not have the same creature comforts of old forests. And if you are a keen angler or simply appreciate a fish-rich stream - you have a parasite to thank for all the fishes.

Sato, T., Watanabe, K., Fukushima, K., & Tokuchi, N. (2014). Parasites and forest chronosequence: Long-term recovery of nematomorph parasites after clear-cut logging. Forest Ecology and Management, 314: 166-171.